Alkaloid polymorphism and ITS sequence variation in the Spiraea japonica complex (Rosaceae) in China: traces of the biological effects of the Himalaya-Tibet Plateau uplift.

Spiraea japonica, a diverse, perennial shrubby species complex widespread across E Asia, was a useful model in a first attempt to link specific phylogeny, floristic evolution, and historical environmental changes in E Asia. DNA-based phylogenetic reconstruction served as a framework to investigate whether the patterns of variation in alkaloids and nrITS from varieties of S. japonica reflect the geological history of China and the uplifting of the Himalaya-Tibet plateau, the most significant geological event in E Asia since the late Paleocene. The high geographical structure found in the phylogenetic trees was corroborated by the distribution of alkaloids. All populations from E China, with mainly hetisine-type alkaloids, formed a single clade, sister to a clade of all populations from SW China, with mainly atisine-type alkaloids. The distribution boundary between the two clades roughly matched the floristic division between the Sino-Japanese Forest and the Sino-Himalayan Forest subkingdoms, as well as the environmental division between SW and E China, suggesting a close link between specific phylogeny, floristic evolution, and geographical changes in E Asia. The divergence between lineages at variety and population level within the eastern clade was slightly older than those within the southwestern clade, supporting the hypothesis of a northeast-southwest migration of Spiraea since the Eocene. The uplift of the Himalaya-Tibet plateau and subsequent increase in geographical complexity in SW China, could facilitate divergence maintenance, thus accelerating the evolutionary rate.